A review of the South African temnospondyl amphibian record R.J. Damiani* & B.S. Rubidge Bernard Price Institute for Palaeontological Research, School of Geosciences, University of the Witwatersrand, Private Bag 3, WITS, Johannesburg, 2050 South Africa Received 14 February 2002. Accepted 30 September 2003 INTRODUCTION The Karoo Basin of South Africa preserves a continuous sequence of sedimentary rocks from the Late Carbonifer- ous through to the Middle Jurassic periods (Smith 1990). Within this thick succession, the Permo-Triassic Beaufort Group and overlying Triassic-Jurassic Stormberg Group preserve a rich fossil biota that documents the evolution of terrestrial life during the late Palaeozoic and early to middle Mesozoic eras. These rocks are world renowned primarily for their therapsid (‘mammal-like reptile’) fauna, the diversity and abundance of which have allowed for the biostratigraphic subdivision of the sequence (Kitching 1977; Kitching & Raath 1984; Rubidge 1995). Less well known amongst Karoo fossil vertebrates are the temnospondyl amphibians, which are surpassed only by therapsids in terms of diversity and abundance. The Temnospondyli was the largest, most diverse and most successful group of fossil amphibians, ranging in time from the Early Carboniferous to the Early Cretaceous (Milner 1990). Temnospondyls were superficially sala- mander-like in appearance with a total body length that varied from below 1 m to 6 m or more, most being above 1 m. They have also been widely linked to the origin of some or all of the modern lissamphibian groups (e.g. Milner 1988; Bolt 1991; Trueb & Cloutier 1991). Temno- spondyls have been recovered from all continents and are near-ubiquitous components of non-marine sedimentary rocks of Permian and Triassic age. This abundance has led to their playing an important and sometimes critical role in local biostratigraphy and global correlation of non- marine strata of Triassic age (e.g. Hancox et al. 1995; Ivakhnenko et al. 1997; Ochev & Shishkin 1989; Lucas 1998). In the Karoo the temnospondyl fauna is dominated by members of the great Mesozoic radiation of temnospondyls known as the Stereospondyli (Milner 1990; Yates & Warren 2000), with members of the Palaeozoic lineages Dvino- sauria and Dissorophoidea making up the non-stereo- spondyl component. The Permo-Triassic Beaufort Group contains all but one of the higher-level temnospondyl taxa known from the Karoo, with most of these occurring in the Triassic part of the Group. The overlying Triassic- Jurassic ‘Stormberg Group’ has received far less attention palaeontologically, a fact reflected in the considerably lower diversity of tetrapods, including temnospondyls (Kitching & Raath 1984). Despite this overall rich temnospondyl record, relatively little research has been directed towards Karoo temnospondyls. In recent years, however, there has been a resurgence of interest in South African temnospondyls, especially in taxonomic studies (see review below), their role in interpreting the basin de- velopment of the Karoo (Hancox & Rubidge 1997; Hancox 1998), and in local biostratigraphy and correlation of non- marine tetrapod faunas (Hancox et al. 1995; Shishkin et al. 1995; Lucas 1998; Damiani 1999; Hancox et al. 2000; Shishkin 2000). The only previous review of Karoo temnospondyls was that of Kitching (1978), a paper which was concerned primarily with documenting the total numbers and strati- graphic distribution of the temnospondyl fauna of the Beaufort Group. Since then, a significant number of new temnospondyl taxa have been described, additional infor- mation has been obtained through redescriptions of older material, and stratigraphic and biostratigraphic advances have been made. The aim of this paper is to summarize the taxonomy as well as stratigraphic occurrence of all known temnospondyls from the Karoo. It is not the intention here to critically reassess the relationships of each temnospondyl species discussed, although we provide a brief summary of the taxonomic history of those species whose relationships have been controversial. In the review below, higher-level taxa are arranged in alphabeti- cal order, whereas individual species within each higher-level taxon are listed in chronological order of description. Synonomy lists include only those references in which a change in a taxon name has occurred. Note that the relationships of the Brachyopidae and Chigutisauridae ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 21 The Karoo of South Africa preserves a near-unbroken sequence of sedimentary rocks and associated vertebrate fauna from the Late Carboniferous to the Middle Jurassic periods. These rocks host a rich temnospondyl amphibian record from the Late Permian to the Early Jurassic. This record is amongst the most diverse temnospondyl record in the world and comprises 10 higher-level taxa, most of which occur in the Triassic part of the sequence and which is dominated by members of the Stereospondyli, with lesser representation by two Palaeozoic lineages. This paper presents a synthesis of the South African temnospondyl amphibian record and highlights the most recent advances in their taxonomy and biostratigraphic placement. Keywords: South Africa, Karoo, temnospondyls, stereospondyls. *Author for correspondence. E-mail: damianir@geosciences.wits.ac.za within the Temnospondyli remain controversial (Shishkin 1991; Schoch & Milner 2000; Yates & Warren 2000; Damiani & Kitching 2003), and are here listed under the Stereospondyli merely for convenience. We precede our review with a brief outline of the Palaeozoic and Mesozoic geology of the Karoo in order to provide the necessary stratigraphic and biostratigraphic framework. Photographs of the skulls of Karoo temnospondyls, in dorsal view, are presented in Figs 1–3. A chart illustrating the biostratigraphic distribution of Beaufort Group temnospondyls is presented elsewhere (Damiani, in press), and is not repeated here. Institutional abbreviations in the text are as follows: AM, Albany Museum, Grahamstown; BMNH, Natural History Museum, London; BP, Bernard Price Institute for Palaeontological Research, Johannesburg; BSP, Bayerische Staatsammlung für Paläontologie und Historische Geologie, Munich; CGP, Council for Geoscience, Pretoria; MA, Museum Africa, Johannesburg; FMNH, Field Museum of Natural History, Chicago; GPIT, Institut und Museum für Geologie und Paläontologie, Tübingen; MGM, MacGregor Museum, Kimberley; NM, National Museum, Bloemfontein; RC, Rubidge Collection, Wellwood, Graaff-Reinet; SAM, South African Museum, Cape Town; TM, Transvaal Museum, Pretoria; UCMP, University of California Museum of Paleontology, Berkeley; UMZC, University Museum of Zoology, Cambridge. GEOLOGICAL SETTING The Karoo Basin of South Africa preserves a 12 km thick succession of mainly sedimentary rocks that accumulated in a retroarc foreland basin during the Late Carboniferous to Early Jurassic in southwestern Gondwana (Johnson et al. 1997; Catuneanu et al. 1998). These rocks reflect changing depositional environments including glacial, deep marine, deltaic, fluvial and aeolian (Smith 1990), and have yielded a diverse fossil biota consisting of plants, vertebrates, invertebrates, and trace fossils (Hancox & Rubidge 1997). Lithostratigraphically the Karoo Supergroup is subdi- vided into five main groups. These are, in ascending stratigraphic order, the Dwyka, Ecca, Beaufort, ‘Stormberg’ and Drakensberg groups, broadly represent- ing deposition in glacial (Dwyka), marine (Ecca) and ter- restrial (Beaufort and Stormberg) environments (Smith 1990). This sedimentary sequence is capped by the Drakensberg volcanics which terminated sedimentation in the basin in the Middle Jurassic (Smith 1990). Vertebrate fossils in this sequence are known from the Ecca, Beaufort and ‘Stormberg’ groups. The Early to Late Permian-aged Ecca Group consists of 16 different formations in the various parts of the basin (Johnson et al. 1997). In terms of vertebrate fossils, actinopterygian fish are common in several of these formations. The mesosaurid reptiles Mesosaurus and Stereosternum are the only tetrapods from the Ecca (Oelofsen 1981, 1987; Modesto 1996), and are confined to the Early Permian Whitehill Formation, a white- weathering, black carbonaceous shale considered to have been deposited in a deep-water, anoxic environment (Oelofsen 1987; Catuneanu et al. 1998). Mesosaurus and Stereosternum are of particular importance as representing the oldest known amniotes from Gondwana. The Beaufort Group conformably overlies the Ecca Group and the lithological contact between these groups is thought to coincide with the position of the ‘palaeo- shoreline’ (Rubidge et al. 2000). Its strata are fluvially derived and typically consist of alternating mudstone and sandstone units with characteristic upward-fining textures, red and purple colours, abundant vertebrate fossils, dessication cracks and palaeosol horizons, suggesting sediment accumulation on vast, semi-arid alluvial plains by floodplain aggradation (Smith 1990). The Beaufort Group preserves a complete Late Permian to early Middle Triassic (Rubidge 1995) sedimentary sequence and associ- ated vertebrate fauna, and has long been recognized as the global standard for the non-marine Permo-Triassic (Romer 1975; Cosgriff 1984; Lucas 1998). This abundant and diverse vertebrate fauna is dominated by therapsids (Kitching 1977) which, combined with the paucity of basin-wide lithostratigraphic markers, have been used for the biostratigraphic subdivision of the group and strati- graphic correlation with faunas elsewhere (Kitching 1977; Keyser & Smith 1979; Rubidge 1995). The current biostratigraphic scheme of the Beaufort Group consists of eight assemblage zones (Rubidge 1995); these have been broadly linked to the lithostratigraphy which comprises many formational names in the various parts of the basin (Keyser & Smith 1979; Rubidge 1995). These biozones are, in ascending order from oldest to youngest, the Eodicynodon, Tapinocephalus, Pristerognathus, Tropidostoma, Cistecephalus, Dicynodon, Lystrosaurus and Cynognathus Assemblage zones. The first six of these biozones are of Late Permian age (Rubidge 1995), the Lystrosaurus Assemblage Zone is broadly Early Triassic in age (Groenewald & Kitching 1995; Hancox 2000), and the Cynognathus Assemblage Zone is of Early-Middle Triassic age (Hancox 2000). Thus contrary to previous opinion which held that the Cynognathus biozone was either of Early (e.g. Anderson & Cruickshank 1978; Cosgriff 1984) or Middle Triassic (e.g. Ochev & Shishkin 1989) age, it is now considered to straddle the Early-Middle Triassic boundary. The Permo-Triassic boundary in the Karoo Basin is currently held to occur within the lowest part of the Lystrosaurus Assemblage Zone, and is defined by the last appearance of the dicynodont Dicynodon (Smith 1995). At present there are no formal subdivisions of the various Beaufort biozones although the Cynognathus Assemblage Zone has been subdivided into a lower ‘A’ zone (upper Olenekian), a middle ‘B’ zone (early Anisian), and an upper ‘C’ zone (late Anisian) (Hancox et al. 1995; Shishkin et al. 1995; Hancox & Rubidge 1997). The ‘Stormberg Group’ unconformably overlies the Beaufort Group and is separated from it by a time gap representing the entire late Middle Triassic (Ladinian) (Hancox 2000). The ‘Stormberg Group’ consists of, in ascending stratigraphic order, the Molteno, Elliot and Clarens formations. The Molteno Formation is of Late Triassic (possibly Carnian) age (Hancox 2000); it hosts a diverse insect and plant assemblage, but as of yet the only 22 ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 vertebrate body fossils are of fish (Anderson & Anderson 1984). The overlying Lower Elliot Formation is regarded by some workers to be a coeval, distal equivalent of the uppermost Molteno Formation (Cairncross et al. 1995; Anderson et al. 1998). It hosts a relatively meagre verte- brate fauna (Anderson et al. 1998) that has been assigned to the Euskelosaurus Range Zone (Kitching & Raath 1984) and used to support a Late Triassic age (Kitching & Raath 1984; Olsen & Galton 1984). The middle and upper parts of the Elliot Formation host a more diverse and abundant vertebrate fauna (Kitching & Raath 1984; Anderson et al. 1998) that, along with the lower part of the overlying Clarens Formation, has been assigned to the Massospondylus Range Zone (Kitching & Raath 1984) and used to support an Early Jurassic age (Smith & Kitching 1997; Anderson et al. 1998). Finally, the Clarens Formation hosts a meagre vertebrate record consisting primarily of elements of the underlying Upper Elliot Formation (Kitching & Raath 1984), and is of Early-Middle Jurassic age. Although vertebrate fossils are considerably less abundant in the Stormberg than in the Beaufort Group, this is compen- sated for by the presence of some of the earliest dino- saurs and mammals in the world (Kitching & Raath 1984). The Elliot Formation also preserves the last known temnospondyls from the Karoo (Warren & Damiani 1999). REVIEW OF SOUTH AFRICAN TEMNOSPONDYLS Temnospondyli Zittel 1887–1890 Dissorophoidea Boulenger 1902 Amphibamidae Moodie 1910 Micropholis stowii Huxley 1859, Fig. 1A Petrophryne granulata Owen 1876a Micropholis granulata Lydekker 1890b Holotype. BMNH R4382, a skull, partially preserved as an internal mould, with attached lower jaw and partial pectoral girdle. Locality and horizon. Given by Huxley (1859) as the ‘foot of the Rhenosterberg’, and by Watson (1913) as ‘Rhenosterberg (north-west of New Bethesda), District Graaff-Reinet, Cape Colony’, although the Rhenosterberg Range lies mainly in the southwest of the adjacent Middelburg District; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. BMNH R510, the holotype skull of ‘Petrophryne granulata’ Owen (Watson 1913); BMNH R510a, a skull (Watson 1913); BSP 1934 VIII 42 and BSP 1934 VIII 43, three and five near-complete skeletons in sandstone nodules, respectively (Broili & Schröder 1937a). All specimens from the farm Donnybrook, Queenstown District, Eastern Cape Province; Lystrosaurus Assemblage Zone, Early Triassic. Remarks. The somewhat enigmatic small temnospondyl Micropholis stowii was initially placed in its own higher- level taxon, the Micropholidae (Watson 1919), which was accepted by most subsequent workers (Haughton 1925; Kuhn 1933; Romer 1947; Huene 1956; Milner 1990; Warren & Hutchinson 1990). However, it is also widely referred to the Dissorophidae (Carroll & Winer 1977; Cosgriff 1984). Most recently, Daly (1994) recognized an assemblage of primitive dissorophoids, including M. stowii, which she transferred to the Amphibamidae. M. stowii is of particular importance as representing the last surviving member of the dissorophoid temnospondyls. The species was redescribed by Boy (1985) on the basis of Broili & Schröder’s (1937a) material, and new material is currently under description by Schoch & Rubidge. Van Heerden (1974) described a supposed skin impression of the reptile Procolophon, which was reinterpreted by Gow (1977) as the cast of a temnospondyl skull roof. Schoch & Milner (2000) attributed that specimen, NM QR1597, to Lydekkerina, but the presence of nodular ornament, large orbits, and a large otic notch (cf. Gow 1977) is identical to M. stowii and unlike Lydekkerina. Dvinosauria Yates & Warren 2000 Tupilakosauridae Kuhn 1960 Thabanchuia oomie Warren 1999, Fig. 1B Holotype. UCMP 42780, a near-complete skull with attached mandibular rami, and associated postcranial elements. Locality and horizon. UCMP locality V4744, near Thaba N’chu, Free State Province; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. UCMP 42777, UCMP 42781, partial skulls and associated postcranial elements, and UCMP 42778, a string of articulated vertebrae with some ribs (Warren 1999); locality and horizon as for holotype. Remarks. Thabanchuia oomie represents the fourth member of the rare higher-level taxon Tupilakosauridae, alongside Tupilakosaurus heilmani from the Early Triassic of East Greenland (Nielson 1954), Tupilakosaurus wetlugensis from the Early Triassic of Russia (Shishkin 1961), and possibly Kourerpeton bradyi from the ?Late Permian of North America (Olsen & Lammers 1976; Warren 1999). The Tupilakosauridae represent relict survivors of the Palaeozoic Dvinosauria (Warren 1999; Yates & Warren 2000). Stereospondyli Zittel 1887–1890 Brachyopidae Lydekker 1885 Batrachosuchus browni Broom 1903, Fig. 1C Holotype. SAM-PK-5868, a near-complete skull. Locality and horizon. An unknown locality in the Aliwal North District, Eastern Cape Province; the horizon is presumably the Cynognathus Assemblage Zone, which is widely exposed in the Aliwal North District. Referred material. None. Remarks. The holotype and only known specimen was collected by Mr Alfred Brown of Aliwal North, and described in an exceedingly brief note by Broom. An attempt was made to trace the type locality from Brown’s extensive personal diaries held at the South Afri- can Museum, but there was nothing on record about its collection. Additional descriptions of Batracho- suchus browni have been provided by Haughton (1925), Watson (1956), Welles & Estes (1969) and Chernin (1977). ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 23 Batrachosuchus sp. Material. UMZC T194, a mandibular fragment, and UMZC T193, an atlas (Watson 1956). UCMP 42856 and UCMP 80859–60, assorted cranial and postcranial remains (Welles & Estes 1969), now renumbered as follows: UCMP 42856, cranial and vertebral remains; UCMP 140568–69, 140571–72, 140575–78 and 140586, skull fragments; UCMP 140587–92, mandibular fragments; UCMP 80857–58, 24 ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 Figure 1. Photographs (in dorsal view) of the skulls of South African temnospondyls. A, Micropholis stowii, SAM-PK-K8550; B, Thabanchuia oomie, holotype, UCMP 42780; C, Batrachosuchus browni, holotype, SAM-PK-5868; D, Bathignathus watsoni, holotype, BMNH R3589; E, Bathignathus poikilops, holotype, BP/1/5790; F, Vanastega plurimidens, holotype, BP/1/4004; G, Laidleria gracilis, holotype, AM 4313; H, Lydekkerina huxleyi, holotype, BMNH R507; I, Eolydekkerina magna, holotype, BP/1/5079. 140580, 140584, 140570, 140573–74, 140583, 140585 and 80859–60, postcranial remains. Locality and horizon. UMZC material from the farm Luiperkop (not ‘Luiper Kop’ of Watson (1956) or ‘Luiperdskop’ of Kitching (1963)), Burgersdorp District, Eastern Cape Province (not ‘Orange Free State’ of Warren & Marsicano (2000)); Cynognathus Assemblage Zone, Early-Middle Triassic. UCMP material from the farm Slootkraal, Rouxville District, Free State Province; Subzone B of the Cynognathus Assemblage Zone, early Anisian. Note that Welles & Estes (1969) ascribe the UCMP material to the farm Bethel; however, the locality from which the material was recovered actually belongs to the farm Slootkraal (Renaut, pers. comm.). Remarks. Further descriptions of some of the UCMP material have been provided by Colbert & Cosgriff (1974) and Warren & Marsicano (2000). Although this material has been referred to Batrachosuchus, there seems no reason why some of this material should not pertain to another brachyopid genus from the Karoo. Bathignathus watsoni (Haughton) Damiani & Jeannot 2002, Fig. 1D Batrachosuchus sp. Watson 1919 Batrachosuchus watsoni Haughton 1925 Holotype. BMNH R3589, a near-complete skull. Locality and horizon. Unknown, but presumed by Watson (1956) to have come from the Burgersdorp District, Eastern Cape Province, and thus from the Cynognathus Assemblage Zone which is widely exposed in this district. Referred material. None. Remarks. Redescribed by Watson (1956) and Welles & Estes (1969), who separated the species from Batrachosuchus browni largely on skull proportions. Chernin (1977) listed further morphological differences and, following Welles & Estes (1969), suggested that ‘Batrachosuchus’ watsoni may be sufficiently distinct from B. browni to warrant generic separation. However, Warren & Marsicano (2000) suggested that the species may be conspecific. Damiani & Jeannot (2002) listed further morphological differences and erected a new genus, Bathignathus, for ‘B.’ watsoni. Bathignathus poikilops Damiani & Jeannot 2002, Fig. 1E Holotype. BP/1/5790, a partial skull roof and part of an occiput. Locality and horizon. Farm Driefontein, Paul Roux District, Free State Province; Subzone A of the Cynognathus Assem- blage Zone, upper Olenekian. Referred material. BP/1/5883, a complete right mandibular ramus (Damiani & Jeannot 2002); locality and horizon as for holotype. Remarks. This brachyopid shows phenetic similarities to both Batrachosuchus browni and Bathignathus (‘Batracho- suchus’) watsoni, but was argued (Damiani & Jeannot 2002) to show closest affinities with the latter. Vanastega plurimidens Damiani & Kitching 2003, Fig. 1F Holotype. BP/1/4004, a partial skull and associated cranial fragments, and a near-complete left mandibular ramus. Paratype. BP/1/5831, a partial left mandibular ramus. Locality and horizon. Farm Nooitgedacht, Burgersdorp District, Eastern Cape Province; Subzone B of the Cynognathus Assemblage Zone, early Anisian. Referred material. None. Remarks. A brachyopid remarkable for apparently being most closely related to the North American Vigilius wellesi (Welles & Estes 1969; Warren & Marsicano 2000), a species suggested (Shishkin 1991; Warren & Marsicano 2000) to be a dvinosaurian. Chigutisauridae Rusconi 1949 cf. Siderops sp. Warren & Damiani 1999 Material. BP/1/5092, a partial skull with attached partial mandibular rami. Locality and horizon. Farm Vastrap, Ladybrand District, Free State Province; Upper Elliot Formation, Early Juras- sic. Remarks. This and the material described below as Chigutisauridae indet. from the Late Triassic-Early Jurassic Elliot Formation represent the last known temnospondyls from the Karoo and the undoubted post-Triassic stereospondyls from Africa. Significantly, all well-known post-Triassic stereospondyl remains from Gondwana appear to be chigutisaurid, including the Elliot material. This material had previously been mentioned by Kitching & Raath (1984), Olsen & Galton (1984) and Hopson (1984), with the Lower Elliot material being referred to as capitosaurid and the Upper Elliot material as brachyopid. All of this material was examined by Warren & Damiani (1999) and found to be chigutisaurid. Chigutisauridae indet. Material. BP/1/4750, fragmentary cranial and postcranial remains from the farm Friesland West, Bethlehem District, Free State Province; Lower Elliot Formation, Late Triassic (Warren & Damiani 1999). BP/1/5111, a partial left mandible with attached skull re- mains, articulated and disarticulated vertebrae, detached partial ribs, and a fragment of interclavicle from the farm Broken Slopes (an annexe of the farm Vastrap), Lady- brand District, Free State Province; Upper Elliot Forma- tion, Early Jurassic (Warren & Damiani 1999). BP/1/5252, partial left and right mandibular rami with associated cranial material from the farm Hollywood, Lady Grey District, Eastern Cape Province; Lower Elliot Formation, Late Triassic (Warren & Damiani 1999). BP/1/5406, fragments of a skull with attached mandibu- lar fragments, vertebral elements and a partial femur from the farm Vastrap, Ladybrand District, Free State Province; Upper Elliot Formation, Early Jurassic (Warren & Damiani 1999). Laidleriidae Kitching 1957 Laidleria gracilis Kitching 1957, Fig. 1G Holotype. AM 4313, an articulated skeleton comprising a complete skull with attached partial lower jaw, and presacral postcranial skeleton. Locality and horizon. Elucwecwe, Engcobo District, East- ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 25 ern Cape Province; Cynognathus Assemblage Zone, Early-Middle Triassic. Referred material. None. Remarks. Kitching (1957) tentatively placed Laidleria gracilis in the Trematosauridae, but suggested that the species may in future be referred to a new ‘family’, the Laidleriidae. The species was retained in the monotypic Laidleriidae by some workers and considered closely related to the Rhytidosteidae (Cosgriff 1965; Warren 1998), but placed directly in the Rhytidosteidae by others (Warren & Black 1985; Milner 1990; Schoch & Milner 2000). Cosgriff & Zawiskie (1979) considered the Laidleriidae as incertae sedis within the Temnospondyli, while Yates & Warren (2000) allied Laidleria with the enig- matic Plagiosauridae. The holotype skull was redescribed in detail by Warren (1998). Lydekkerinidae Watson 1919 Lydekkerina huxleyi (Lydekker) Broom 1915, Fig. 1H Bothriceps huxleyi Lydekker 1889 Putterillia platyceps Broom 1930 Limnoiketes paludinatans Parrington 1948 Lydekkerina sp. Parrington 1948 Holotype. BMNH R507, a complete skull. Locality and horizon. An unknown locality near Eden- burg, Free State Province; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. BMNH R504–506, R508, cranial and postcranial remains in sandstone nodules (Lydekker 1890b); locality and horizon as for holotype. SAM- PK-3525, a partial skull (Haughton 1925); BSP 1934 VIII 44, three skulls with associated partial postcranial skeletons in a sandstone nodule (Broili & Schröder 1937b); UMZC T238, a skull and associated cervical vertebrae (Parrington 1948); a partial skull and associated postcrania described as Lydekkerina sp. by Parrington (1948); UMZC T214, the holotype (a skull and pectoral girdle) of Limnoiketes paludinatans Parrington (Shishkin et al. 1996); TM 168, TM 88, the holotype (a skull) and paratype (a dermal shoulder girdle and lower jaw), respectively, of Putterillia platyceps Broom (Shishkin et al. 1996). All material except for BMNH specimens are from the Harrismith Commonage, Free State Province; Lystrosaurus Assemblage Zone, Early Tri- assic. Remarks. Watson (1912, 1919) provided a redescription of some of the British Museum material, while Shishkin et al. (1996) provide additional data based on specimens from the TM, BPI and NM. Despite a wealth of specimens, the ontogeny and morphological variation within Lydekkerina huxleyi has never been studied, and forms the basis of a current research project by A.M. Jeannot. This study will also determine if only one species of Lydekkerina is present, as is universally assumed. Broomulus dutoiti (Broom) Romer 1947 Lydekkerina dutoiti Broom 1930 Holotype. MGM 4285, a poorly preserved skull with associated lower jaws and partial pectoral girdle. Locality and horizon. Harrismith Commonage, Free State Province; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. None. Remarks. Variously considered a valid species, indeter- minate, or a junior synonym of Lydekkerina huxleyi. The species was redescribed following additional preparation by Shishkin et al. (1996) who confirmed its status as a valid taxon differing from L. huxleyi by a suite of characters. Eolydekkerina magna Shishkin et al. 1996, Fig. 1I Holotype. BP/1/5079, a near-complete skull. Locality and horizon. Farm Fairydale, Bethulie District, Free State Province; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. None. Mastodonsauridae Lydekker 1885 sensu Damiani 2001a Xenotosuchus africanus (Broom) Morales & Shishkin 2002, Fig. 2A Capitosaurus africanus Broom 1909 nomen nudum Capitosaurus africanus (Broom) Haughton 1925 Parotosaurus africanus (Broom) Watson 1962 Parotosuchus africanus (Broom) Chernin 1978 Wellesaurus africanus (Broom) Damiani 2001a Lectotype. SAM-PK-2360, the posterior half of a skull and a partial left mandible; designated type by Haughton (1925), since Broom (1909) did not designate a holotype. Locality and horizon. Farm Vaalbank, Burgersdorp District, Eastern Cape Province; Subzone B of the Cynognathus Assemblage Zone, early Anisian. Referred material. SAM-PK-3008, a poorly preserved skull (Haughton 1925); farm Winaarsbaken, Burgersdorp District, Eastern Cape Province. UCMP 41286, a near- complete skull and lower jaws (Morales & Shishkin 2002); farm Matabele, Rouxville District, Free State Province. Both specimens from Subzone B of the Cynognathus Assemblage Zone; early Anisian. Remarks. Welles & Cosgriff (1965) considered ‘Capito- saurus’ africanus a nomen ‘vanum’ since Broom (1909) did not figure the skull and provided only a cursory description. Chernin (1978) figured and redescribed the lectotype and transferred the species to the catch-all genus Parotosuchus. Shishkin et al. (1995), Maryanska & Shishkin (1996) and Damiani (2001a) suggested that ‘Parotosuchus’ africanus may be related to Wellesaurus peabodyi (Welles & Cosgriff 1965; Damiani 2001a), and Damiani (2001a) transferred ‘P.’ africanus to Wellesaurus. Most recently, Morales & Shishkin (2002) described a new specimen of W. africanus and transferred the species to a new genus, Xenotosuchus, but conceded that the species showed closest affinities to ‘stenotosaurids’ (including Wellesaurus). Parotosuchus haughtoni (Broili & Schröder) Chernin 1978, Fig. 2B Capitosaurus haughtoni Broili & Schröder 1937c Parotosaurus haughtoni (Broili & Schröder) Romer 1947 Karoosuchus haughtoni (Broili & Schröder) Ochev 1966 Holotype. BSP 1934 VIII 50, a fragmentary skull, apparently destroyed during World War II. Neotype: 26 ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 BP/1/5673, a partial skull and lower jaws, and associated intercentra (Damiani 2001b), designated neotype by Damiani (2001a). Locality and horizon. Holotype: farm Kaaimansgat, Rouxville District, Free State Province; Subzone A of the Cynognathus Assemblage Zone, upper Olenekian. Neo- type: Farm Driefontein, Paul Roux District, Free State Prov- ince; Subzone A of the Cynognathus Assemblage Zone, upper Olenekian. Referred material. None. Remarks. This species was long known from the holo- type specimen only and was usually referred by most ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 27 Figure 2. Photographs (in dorsal view) of the skulls of South African temnospondyls. A, Xenotosuchus africanus, holotype, SAM-PK-2360; B, Parotosuchus haughtoni, neotype, BP/1/5673; C, Watsonisuchus magnus, holotype, UMZC T173; D, Paracyclotosaurus morganorum, holotype, BP/1/5551; E, Jammerbergia formops, holotype, NM QR1436; F, Uranocentrodon senekalensis, referred specimen, NM QR1483; G, Laccosaurus watsoni, referred specimen, BP/1/213; H, Broomistega putterilli, referred specimen, BP/1/5058; I, Rhinesuchoides tenuiceps, holotype, FMNH UC 1519. workers to the catch-all genus Parotosuchus (=‘Capitosaurus’ and ‘Parotosaurus’), but placed in a new genus, Karoosuchus, by Ochev (1966). However, Damiani (1999, 2001b) described a new, near-complete specimen of Parotosuchus haughtoni which was made the neotype for the species (Damiani 2001a). Watsonisuchus magnus (Watson) Ochev 1966, Fig. 2C Wetlugasaurus sp. Watson 1919 Wetlugasaurus magnus Watson 1962 Parotosaurus sp. Welles & Cosgriff 1965 Parotosuchus sp. Warren 1980 Holotype. UMZC T173, the posterior half of a skull split into several sections, and fragments of a right mandible. Locality and horizon. Farm Watford, Burgersdorp District, Eastern Cape Province; Subzone A of the Cynognathus Assemblage Zone, upper Olenekian. Referred material. None. Remarks. Considered by Watson (1919, 1962) to most closely resemble the Laurasian taxon Wetlugasaurus in general skull morphology, later transferred to the catch-all taxon ‘Parotosaurus’ by Welles & Cosgriff (1965), but shown by Ochev (1966) to pertain neither to Wetlugasaurus or ‘Parotosaurus’ (=Parotosuchus). Watsonisuchus sp. Damiani et al. 2001 Material. CGP 1/60, the posterior portion of a right mandible. Locality and horizon. Farm T’Zamenkomst 120, Venter- stad District, Eastern Cape Province; Lystrosaurus Assem- blage Zone, Early Triassic. Remarks. This specimen represents the first undoubted record of a mastodonsaurid from the Lystrosaurus biozone. Paracyclotosaurus morganorum Damiani & Hancox 2003, Fig. 2D Parotosuchus morgani Hancox & Rubidge 1997, nomen nudum Paracyclotosaurus Hancox et al. 2000 Holotype. BP/1/5551, the right posterior portion of a skull. Locality and horizon. Farm Wilgerkloof, Sterkstroom District, Eastern Cape Province; Subzone C of the Cynognathus Assemblage Zone, late Anisian. Referred material. None. Jammerbergia formops Damiani & Hancox 2003, Fig. 2E Holotype. NM QR1436, a partial skull. Locality and horizon. Jammerberg, Wepener District, Free State Province; the horizon is presumably the Cynognathus Assemblage Zone (see discussion in Damiani & Hancox 2003). Referred material. None. Mastodonsauridae indet. Cyclotosaurus albertyni Broom 1904, nomen dubium Holotype. SAM-PK-K1876, fragments of a skull roof. Locality and horizon. An unknown locality in the Rouxville District, Free State Province; the horizon is presumably the Cynognathus Assemblage Zone, which is widely exposed in the Rouxville District. Referred material. None. Remarks. Broom (1904) stated that the frontal was included in the orbital margin and considered the otic notch to have been closed posteriorly, as in the Late Triassic Cyclotosaurus. The specimen is likely to be mastodonsaurid but the exceedingly brief description and lack of figures renders any taxonomic assignment uncertain. The holotype could not be located in the SAM collections and is evidently lost. Kestrosaurus dreyeri Haughton 1925 Holotype. SAM-PK-3452, a fragmentary skull, the fragments of which are set in plaster. Locality and horizon. Farm Harmonia, Senekal District, Free State Province; Subzone A of the Cynognathus Assem- blage Zone, upper Olenekian. Referred material. A mandibular fragment said to be associated with the holotype skull (Chernin 1978). Remarks. Kestrosaurus dreyeri was initially described (Haughton 1925) as possessing, amongst other peculiari- ties, paired anterior palatal vacuities, circular nostrils and an interfrontal. The presence of these characters was apparently corroborated by Chernin (1978), who redescribed the holotype skull as well as a referred mandibular fragment. K. dreyeri has been variously considered an aberrant ‘capitosaurid’ (Welles & Cosgriff 1965; Chernin 1978), closely related to Benthosuchus (Romer 1947), or closely related to Mastodonsaurus (Shishkin 1980; Milner et al. 1990), primarily on the basis of the alleged presence of paired anterior palatal vacuities. More recent workers (Hancox et al. 1995; Shishkin et al. 1995) consider K. dreyeri a typical ‘capitosaurid’ closely related to Parotosuchus. However, Damiani (2001a) considered the holotype skull an indeterminate masto- donsaurid as it was based largely on a plaster reconstruc- tion within which non-diagnostic mastodonsaurid fragments had been embedded. Parotosuchus dirus Chernin 1978 Holotype. SAM-PK-434, fragments of a left and right lower jaw and associated cranial fragments. Locality and horizon. An unknown locality near Aliwal North, Eastern Cape Province; the horizon is presumably the Cynognathus Assemblage Zone, which is widely exposed in the Aliwal North District. Referred material. UMZC T167, a mastodonsaurid fragment from Aliwal North, Eastern Cape Province (Schoch & Milner 2000). Remarks. The holotype pertains to a very large individ- ual and is unusual in showing an additional row of teeth posterior to the usual post-symphyseal tooth row, and teeth on the anterior coronoid. The latter character is rare in stereospondyls but does occur in undescribed mandi- bles from the Cynognathus Assemblage Zone, probably referable to Watsonisuchus (RJD, pers. obs.). However, those mandibles are considerably smaller than that of P. dirus, differ significantly in the structure of the post-glenoid area, and usually lack any parasymphyseal teeth (RJD, pers. obs.). Parotosuchus dirus does not pertain 28 ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 to Parotosuchus and probably represents a new genus, but additional material is required for a definitive diagnosis. Parotosaurus sp. Material. UMZC T135, the posterior portion of a left mandible (Watson 1962). Locality and horizon. Farm Winaarsbaken, Burgersdorp District (not ‘Prince Albert District’ of Schoch & Milner (2000)), Eastern Cape Province; Subzone B of the Cyno- gnathus Assemblage Zone, early Anisian. Remarks. As the crucial post-glenoid area of the mandi- ble is damaged the specimen is largely generically inde- terminate. However, it likely pertains to Wellesaurus africanus which is known from this locality and is the most common mastodonsaurid in Subzone B of the Cynognathus biozone. Rhinesuchidae Watson 1919 Rhinesuchus whaitsi Broom 1908 Rhinesuchus beaufortensis Boonstra 1940 Holotype. SAM-PK-1212, the basicranial region of a skull and associated cranial fragments. Locality and horizon. Near Leeu-Gamka (formerly Fraserburg Road Station), Prince Albert District, Western Cape Province; Tapinocephalus Assemblage Zone, Late Permian. Referred material. SAM-PK-3009, a near-complete skull and partial lower jaws (Haughton 1915b) from an unknown locality near Beaufort West, Western Cape Province. Skull fragments (unnumbered) from the farm Zeekoegat, Prince Albert District, Western Cape Province, mentioned by Broom (1908, 1912); Tapinocephalus Assem- blage Zone, Late Permian. The posterior half of a skull from the farm Blaauw Krantz, Prince Albert District, Western Cape Province, mentioned by Haughton (1925); Tapinocephalus Assemblage Zone, Late Permian. SAM-PK-9135, the posterior end of a mandible (Watson 1962); farm Vogelfontein, Prince Albert District, Tapinocephalus Assemblage Zone, Late Permian. UMZC T64, a basicranial fragment (Watson 1962); farm Zeekoegat, Prince Albert District, Western Cape Province; Tapinocephalus Assemblage Zone, Late Permian. Remarks. The Rhinesuchidae require a thorough morphological revision as inadequate data are available in the literature and most of the material is in need of prepa- ration (RJD, pers. obs.). Consequently, the taxonomy and all accompanying remarks in this section are necessarily provisional pending a forthcoming revision by the authors. Haughton (1915b) briefly described a skull (SAM-PK- 3009) which he referred to R. whaitsi but provided only photographs of the specimen and drawings of cross- sections through the left mandible. Watson (1919) figured the palate of R. whaitsi based on a sketch sent to him by Broom, and which Watson (1962) later considered to have probably been based on SAM-PK-3009. Haughton (1925) subsequently redescribed the holotype basi- cranial fragment and designated SAM-PK-3009 the ‘para- type’ for the species; however, this designation cannot hold as Broom (1908) did not designate a paratype. SAM-PK-3009 was later designated the holotype for the new species Rhinesuchus beaufortensis (Boonstra 1940), with which Watson (1962) agreed but which is here and in Schoch & Milner (2000) considered a junior synonym of R. whaitsi. Uranocentrodon senekalensis van Hoepen 1917, Fig. 2F Myriodon senekalensis van Hoepen 1911 Rhinesuchus major Broom 1912 Rhinesuchus senekalensis (van Hoepen) Haughton 1915b Holotype. TM 75, the right side of a partial skull with attached lower jaw (van Hoepen 1911). Locality and horizon. A sandstone quarry, the precise locality of which remains uncertain, near Senekal, Free State Province; Dicynodon Assemblage Zone, Late Permian. Referred material. MA 60c1 and 60c1a, two partial skulls, described by Broom (1912) as Rhinesuchus major, and men- tioned by van Hoepen (1911, 1915) as the specimens that had ‘previously found their way to the Museum of the University College at Johannesburg’; three near-complete skeletons in numerous blocks of sandstone, all originally in the Transvaal Museum, Pretoria, and with which the holotype specimen, TM 75, is associated (van Hoepen 1915); NM QR1483, a near-complete skeleton in a sand- stone block (Haughton 1915b); SAM-PK-2783, a partial left mandible with attached pieces of skull mentioned by Haughton (1915b); TM 185, a near-complete skull with attached lower jaw (Broom 1930). All known material of Uranocentrodon senekalensis is presumed to have come from the same unknown quarry near Senekal, a hypothe- sis supported by the similar lithology of the matrix associated with each specimen (Latimer et al. 2002). Remarks. Uranocentrodon senekalensis was long consid- ered from the Early Triassic Lystrosaurus Assemblage Zone (e.g. Kitching 1978) but has recently been shown (Latimer et al. 2002) to belong to the underlying Dicynodon Assem- blage Zone, the assumption being that all of the known material was recovered from the same quarry near Senekal. In addition to the principal descriptions given by van Hoepen, Broom and Haughton, additional data on the species is provided by Watson (1962) and Findlay (1968). Laccosaurus watsoni Haughton 1925, Fig. 2G Lydekkerina kitchingi Broom 1950 Muchocephalus muchos Watson 1962 Uranocentrodon watsoni (Haughton) Ochev 1966 Muchocephalus kitchingi (Broom) Schoch & Milner 2000 Holotype. SAM-PK-4010, a near-complete skull. Locality and horizon. Farm Ferndale, Graaff-Reinet District, Eastern Cape Province; Dicynodon (formerly Daptocephalus) Assemblage Zone, Late Permian (Kitching 1978). Referred material. BP/1/215, the holotype (a near- complete skull) of Lydekkerina kitchingi Broom. BP/1/213, the holotype (a near-complete skull) of Muchocephalus muchos Watson. Both specimens were found in close association on the farm Ringsfontein, Murraysburg District (Kitching 1978); Dicynodon (formerly Dapto- cephalus) Assemblage Zone, Late Permian. An additional ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 29 92 specimens (reposited in the BPI, Johannesburg) from the same horizon on the adjoining farm Beeldhouersfontein were referred to L. kitchingi by Kitching (1977, 1978). Remarks. It appears likely that the holotypes of Lydekkerina kitchingi (Broom 1950), which is based on a small skull, and Muchocephalus muchos (Watson 1962), a much larger skull, represent juvenile and adult individu- als of the same species as they were recovered in close association from the same locality. As the holotype of M. muchos appears morphologically indistinguishable from that of Laccosaurus watsoni (Haughton 1925), both L. kitchingi and M. muchos are here considered subjective junior synonyms of L. watsoni. As far as can be determined the 92 additional specimens of L. kitchingi reported by Kitching (1977, 1978) consist exclusively of small, probably juvenile, skulls (RJD, pers. obs.). Kitching (1978) also mentions a paratype specimen of L. kitchingi from Ringsfontein; however, Broom (1950) did not designate a paratype. Rhinesuchus capensis Haughton 1925 Rhinesuchus avenanti Boonstra 1940 Holotype. SAM-PK-7419, an incomplete skull. Locality and horizon. Farm Spitzkop, Graaff-Reinet District, Eastern Cape Province; the horizon was given by Haughton (1925) as the ‘Endothiodon zone’ (now corre- sponding to the Tropidostoma Assemblage Zone), and by Broom (1948) and Kitching (1978) as the ‘Cistecephalus zone’, the latter now corresponding to both the Tropidostoma and Cistecephalus Assemblage zones (Rubidge 1995). Referred material. SAM-PK-11489, the holotype (a skull) of Rhinesuchus avenanti Boonstra; farm Mynhardts Kraal, Beaufort West District, Western Cape Province. The hori- zon is stated as the ‘Tapinocephalus zone’ by Boonstra (1940) and Kitching (1977), which now corresponds to both the Tapinocephalus and Pristerognathus Assemblage zones. Remarks. The holotype of R. capensis is the largest known rhinesuchid skull with a high degree of ossification of the cranial bones (Haughton 1925). Consequently, sutures are poorly shown. Note that figures of R. capensis given by Haughton (1925) are inaccurate in that the skull is depicted as too short and broad, whereas it should be somewhat more slender and elongated (RJD, pers. obs.). The holotype of R. avenanti is considerably smaller but is otherwise very similar morphologically to R. capensis, and is here considered a junior synonym of the latter. In contrast, Watson (1962) and Schoch & Milner (2000) considered R. avenanti a junior synonym of Rhinesuchoides tenuiceps (Olson & Broom 1937), probably on the basis of having a similarly slender skull. Broomistega putterilli (Broom) Shishkin & Rubidge 2000, Fig. 2H Lydekkerina putterilli Broom 1930 Holotype. TM 184, the posterior half of a skull and an associated mandibular fragment. Locality and horizon. Harrismith Commonage, Free State Province; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. BP/1/5058, a complete skull with lower jaw in articulation, and associated dermal pectoral girdle (Shishkin & Rubidge 2000); Ndanyane Hill, farm Hartebeest Spruit, Bergville District, KwaZulu-Natal. BP/1/3241, a partially disarticulated skeleton including skull and lower jaw (Shishkin & Rubidge 2000); farm Admiralty Estates, near Oliviershoek Pass, Bergville District, KwaZulu-Natal. Both specimens from the Lystrosaurus Assemblage Zone, Early Triassic. Remarks. Described originally by Broom (1930) as a lydekkerinid closely related to Lydekkerina huxleyi, but shown by Shishkin & Rubidge (2000) to be a rhinesuchid and the only one known from the Triassic. Rhinesuchoides tenuiceps Olson & Broom 1937, Fig. 2I Rhinesuchus rubidgei Broom 1948 Holotype. FMNH UC 1519, a skull and partial left lower jaw. Locality and horizon. Stated by Olson & Broom (1937) as ‘Tapinocephalus zone, 2 miles east of Stinkfontein’; this is probably the farm Stinkfontein located in the Prince Albert District, Western Cape Province, in which the Tapinocephalus Assemblage Zone is exposed (Kitching 1977). Referred material. RC 73, the holotype (a skull) of Rhinesuchus rubidgei Broom (Schoch & Milner 2000); Farm Wimbledon (Welgevonden), Graaff-Reinet District, Eastern Cape Province; Cistecephalus Assemblage Zone, Late Permian. Remarks. The synonomy of Rhinesuchus rubidgei and Rhinesuchoides tenuiceps was also accepted by Schoch & Milner (2000), both specimens being based on similarly sized skulls with near-identical proportions. Schoch & Milner (2000) also referred to R. tenuiceps three specimens, BP/1/2741, BP/1/2931 and BP/1/2958, which they state as pertaining to the Tapinocephalus Assemblage Zone. However, the latter two specimens are small rhinesuchid skulls associated with those collected by Kitching on the farm Beeldhouersfontein in the Murraysburg District (Kitching (1977, 1978), which were considered to belong to ‘Lydekkerina’ kitchingi (see Laccosaurus watsoni) and are from the Dicynodon Assemblage Zone. BP/1/2741 is a partial pelvis of the therapsid Lystrosaurus from the Lystrosaurus Assemblage Zone; the number as stated is probably a typographical error. Rhinesuchidae indet. Eryops oweni Lydekker 1890a, nomen dubium Eryops africanus Lydekker 1890b Rhinesuchus africanus (Lydekker) Broom 1908 Holotype. BMNH R466, an incomplete right mandibular ramus. Locality and horizon. An unknown locality in the Karoo. Referred material. BMNH R470, an intercentrum (Lydekker 1890a); locality and horizon unknown but apparently associated with the holotype. SAM-PK-3010, a fragmentary skull and lower jaw (Haughton 1915b); farm Dunedin, Beaufort West District, Western Cape Province; the horizon was stated by Haughton (1915b) as ‘Ciste- cephalus zone’, although Kitching (1977) stated that the 30 ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 locality pertains to the former ‘Endothiodon zone’, which now corresponds to the Tropidostoma Assemblage Zone (Rubidge 1995). Remarks. The holotype specimen almost certainly pertains to a rhinesuchid because of the dense shagreen field covering the coronoid series and the slightly antero-posteriorly compressed marginal teeth. Haughton’s (1915b) referred specimen includes a fragmentary and badly damaged skull and mandible; however, there is no reason to consider this specimen conspecific with Lydekker’s holotype and the state of preservation of SAM-PK-3010 precludes any taxonomic referral. Rhinesuchus broomianus von Huene 1931, nomen dubium Holotype. GPIT Huene 1931 F.1, a partial skull. Locality and horizon. Farm Kuilspoort, Beaufort West District, Western Cape Province; Tropidostoma (previously Endothiodon) Assemblage Zone, Late Permian. Referred material. None. Remarks. This species was considered valid by Schoch & Milner (2000) whose diagnosis was based principally on the skull being somewhat broader than in other rhinesuchids. However, the skull of R. broomianus has evidently undergone considerable dorso-ventral compac- tion and is poorly preserved, rendering any comparisons with other rhinesuchids difficult. Rhytidosteidae Huene 1920 Rhytidosteus capensis Owen 1884 Holotype. BMNH R455, the antorbital portion of a skull with the anterior parts of the lower jaw in articulation, and the posterior part of the right mandible. Locality and horizon. Given by Owen as ‘Beersheba’, Free State Province; Horizon unknown because of ambiguous locality details. Considered variably as from either the Lystrosaurus (Kitching 1978; Cosgriff & Zawiskie 1979; Shishkin 1994) or Cynognathus (Huene 1920; Romer 1947; Warren & Black 1985) Assemblage Zones. Referred material. BMNH R503, an interotic fragment, a basicranial fragment, and a clavicular fragment (Lydekker 1890b), all considered by Lydekker to have been associ- ated with the holotype; locality and horizon as for the holotype. Remarks. Additional descriptions of Rhytidosteus capensis have been provided by Watson (1919), Huene (1920), Romer (1947) and Cosgriff (1965). Romer (1947) consid- ered R. capensis conspecific with the trematosaurid Microposaurus casei (Haughton 1925), reconstructing the former using the outline of the latter and transferring R. capensis to the Trematosauridae. Cosgriff (1965) considered the association of R. capensis with M. casei to have been erroneous and returned the former to the Rhytidosteidae. Warren & Black (1985) suggested that R. capensis may be a composite of rhytidosteid and trematosaurid fragments, and left the species incertae sedis within the Trematosauroidea (sensu Warren & Black 1985: Rhytidosteidae + Trematosauridae). However, most workers (Cosgriff 1984; Milner 1990; Shishkin 1994; Schoch & Milner 2000) consider R. capensis a valid rhytidosteid. Schoch & Milner (2000) listed the fragmen- tary Russian species Rhytidosteus uralensis (Shishkin 1994) as a junior synonym of R. capensis. Although this assump- tion is reasonable, we prefer to leave R. uralensis as a separate species pending more complete material. Pneumatostega potamia Cosgriff & Zawiskie 1979, Fig. 3A Holotype. BP/1/981, a sandstone fragment containing the impression of a partial skull roof. Locality and horizon. Farm Tafelberg, Middelburg District, Eastern Cape Province; Lystrosaurus Assemblage Zone, Early Triassic. Referred material. SAM-PK-11188, associated fragments of a skull, lower jaw, dermal shoulder girdle and vertebral column (Cosgriff & Zawiskie 1979); farm Goede Hoop, Colesberg District, Eastern Cape Province; Lystrosaurus Assemblage Zone, Early Triassic. Trematosauridae Watson 1919 Trematosuchus sobeyi (Haughton) Watson 1919, Fig. 3B Trematosaurus sobeyi Haughton 1915a Aphaneramma sp. Hammer 1987, in part. Holotype. SAM-PK-2779, a complete skull. Locality and horizon. Sobey’s Quarry, Queenstown, Eastern Cape Province; Subzone A of the Cynognathus Assemblage Zone, upper Olenekian. Referred material. SAM-PK-5136, the middle portion of a skull (Haughton 1925); locality and horizon as for the holotype. NM QR3263A and B, the anterior part of a snout and a palatal fragment, respectively (Shishkin & Welman 1994); farm Verdun, Paul Roux District, Free State Province; Subzone A of the Cynognathus Assemblage Zone, upper Olenekian. Remarks. The referred specimen, SAM-PK-5136, was considered an indeterminate long-snouted (i.e. loncho- rhynchine) trematosaurid and thus transferred to Aphane- ramma sp. by Hammer (1987). However, SAM-PK-5136 comes from the same locality as the holotype of Trematosuchus sobeyi and is therefore reasonably consid- ered referable to that taxon. The difficulty in placing SAM-PK-5136 taxonomically stems from the morphologi- cal similarity in postorbital skull morphology between short and long-snouted trematosaurids. A redescription of SAM-PK-5136 and all other supposed lonchorhyn- chines from the Karoo is in progress by the senior author. Microposaurus casei Haughton 1925, Fig. 3C Rhytidosteus casei Shishkin 1964 Holotype. SAM-PK-6556, a complete skull. Locality and horizon. Exposures just above the Wonderboom Bridge, Burgersdorp District, Eastern Cape Province; Subzone B of the Cynognathus Assemblage Zone, early Anisian. Referred material. None. Remarks. Considered by Romer (1947) and Shishkin (1964) to be congeneric with the fragmentary rhytidosteid Rhytidosteus capensis, but shown by Cosgriff (1965) to be a trematosaurid. The species was later excluded from the Trematosauridae by Hammer (1987), but included by ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 31 Warren & Black (1985), Hellrung (1987) and Schoch & Milner (2000). The latter authors suggested close affinities with Tertrema from the Early Triassic of Spitzbergen. However, the morphology of M. casei, currently being redescribed following additional preparation by the senior author, points strongly to affinities with the Russian ‘platystegid’ Inflectosaurus amplus (Shishkin 1960). Trematosauridae indet. Trematosaurus kannemeyeri Broom 1909, nomen dubium Aphaneramma kannemeyeri (Broom) Huene 1920 Trematosuchus kannemeyeri (Broom) Zittel 1923 Gonioglyptus kannemeyeri (Broom) Huene 1925 Aphaneramma sp. Hammer 1987, in part Holotype. SAM-PK-1329, a portion of the middle part of a badly weathered skull, including the orbits. Locality and horizon. An unknown locality in the Free State Province, then known as the ‘Orange River Colony’ (Broom 1909); horizon unknown, but suggested as ‘Upper Beaufort Beds’ (equivalent to the Cynognathus Assem- blage Zone) by Haughton (1925), and adopted by all subsequent workers. Referred material. None. Remarks. Widely considered an indeterminate, Aphaneramma-like lonchorhynchine by recent workers (Hammer 1987; Welles 1993; Shishkin & Welman 1994), and listed as a valid lonchorhynchine, Aphaneramma kannemeyeri, by Schoch & Milner (2000). In contrast, Damiani et al. (2000) considered Trematosaurus kannemeyeri referrable to either a long or short-snouted trematosaurid, and declared the specimen a nomen dubium. Subsequent re-examination by the senior author has confirmed its lonchorhynchine affinities; however, its poor state of preservation precludes any further taxonomic placement and the species should remain a nomen dubium. A redescription is currently in progress by the senior author. A recently discovered lonchorhynchine from the Lystrosaurus Assemblage Zone was reported by Damiani & Welman (2001) as representing an undoubted lonchorhynchine from that biozone. This specimen, NM QR3424, is represented by a partial snout and may belong to the same species as ‘Trematosaurus’ kannemeyeri. A full description will be the subject of a forthcoming publica- tion. Trematosauridae gen. et. sp. indet. Material. CGP 1/5, a complete but damaged right mandibular ramus (Damiani et al. 2000). Locality and horizon. Farm Perdelaagte (an annexe of the farm Heuningkrans 26), Burgersdorp District, Eastern Cape Province; Lystrosaurus Assemblage Zone, Early Triassic. Remarks. This specimen appears to belong to a short- snouted trematosaurid such as Trematosaurus or Tremato- suchus, and, along with NM QR3424, represents the sec- ond trematosaurid taxon from the Lystrosaurus Assem- blage Zone. The specimen is of further importance as it comes from high in the Lystrosaurus Assemblage Zone, in strata which have been postulated to pertain to an ‘inter- mediate’ biozone between the Lystrosaurus and Cynognathus biozones (Neveling et al. 1999). Stereospondyli indet. Laccocephalus insperatus Watson 1919, nomen dubium Holotype. BMNH R532, an incomplete skull in a sandstone nodule. Locality and horizon. Stated by Watson (1919) as ‘near Mr Hope’s farm, Orange Free State’; horizon unknown. Referred material. None. Remarks. The poor preservation of L. insperatus renders any taxonomic assignment uncertain. However, in size and general outline the specimen is almost certainly a 32 ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 Figure 3. Photographs (in dorsal view) of the skulls of South African temnospondyls. A, Pneumatostega potamia, holotype, BP/1/981; B, Trematosuchus sobeyi, holotype, SAM-PK-2779; C, Microposaurus casei, holotype, SAM-PK-6556. stereospondyl. In particular, the presence of a crista muscularis of the parasphenoid and a median, raised area on the cultriform process, as alluded to by Watson (1919), suggest that L. insperatus may be a rhinesuchid or mastodonsaurid. However, there is neither sufficient morphological nor stratigraphic data to suggest that L. insperatus is synonymous with Uranocentrodon senekalensis (contra Schoch & Milner 2000). Stereospondyli gen. et. sp. indet. Material. BP/1/4935, a fragment of a mandibular ramus from the farm Mequatling, Clocolan District, Free State Province (Warren & Damiani 1999); Lower Elliot Forma- tion, Late Triassic. Remarks. Likely to pertain to a chigutisaurid, as with all of the other known temnospondyl material from the Elliot Formation (see Chigutisauridae). Temnospondyli indet. Petrophryne major Owen 1876b, nomen dubium Bothriceps major (Owen) Lydekker 1890b Holotype. BMNH R4145, a sandstone block containing a poorly preserved partial skull, largely as impression. Originally held in the museum of the Geological Society of London under the catalogue number S.A.119. Locality and horizon. Collection data associated with the holotype states the horizon and locality as the ‘Stormberg Beds’, ‘Southern margin of the Stormberg Range’. Ptychosphenodon browni Seeley 1907, nomen dubium Holotype. BMNH R3503, a mandibular fragment. Locality and horizon. An unknown locality near Aliwal North, Eastern Cape Province; the horizon is presumably the Cynognathus Assemblage Zone, which is widely exposed in the Aliwal North District. Syphonodon thecomastodon Seeley 1908, nomen dubium Holotype. BMNH R3609, a large temnospondyl tooth. Locality and horizon. Farm Wonderboom, Burgersdorp District, Eastern Cape Province; the horizon is presum- ably the Cynognathus Assemblage Zone, which is widely exposed in the Burgersdorp District. Phrynosuchus whaitsi Broom 1913, nomen dubium Rhinesuchus sp. Chernin & Kitching 1977 Holotype. SAM-PK-2357, a crushed, poorly preserved skull and articulated partial postcranial skeleton in a sand- stone nodule. Locality and horizon. Farm Droogvoets Fontein, Fraser- burg District, Western Cape Province; Horizon uncertain, but suggested by Broom (1913) as ‘upper part of the Endothiodon Zone’, and more recently (Chernin & Kitching 1977) as either uppermost Tapinocephalus Zone or lowermost Cistecephalus Zone. These horizons now correspond, respectively, to the Pristerognathus and Tropidostoma Assemblage Zones, of Late Permian age. Remarks. Broom (1913) made no attempt to place the specimen taxonomically, but it was later considered an indeterminate rhinesuchid by Chernin & Kitching (1977). Schoch & Milner (2000) followed this assignation. How- ever, re-examination of the specimen by the senior author failed to reveal any resemblance to rhinesuchids or any characters which could place the specimen taxonomically. CONCLUSIONS The Karoo of South Africa boasts a rich and varied temnospondyl fauna that is amongst the most diverse in the world, with 10 higher-level taxa recorded to date. This record is dominated by predominantly large temno- spondyls of the clade Stereospondyli. The Palaeozoic temnospondyl record in the Karoo is at present restricted to one higher-level taxon, the Rhinesuchidae, despite the fact that six of the eight biozones of the Beaufort Group are of Permian age. It is unclear, however, whether this reflects a true paucity (or absence) of other higher-level taxa or a preservational bias (Kitching 1978). Members of the Rhinesuchidae appear to be endemic to Gondwana, and have been recorded else- where from the Upper Permian of what is now Malawi (Watson 1962), India (Werneburg & Schneider 1996) and Brazil (Barberena 1998; Barberena & Dias 1998). However, the diversity of rhinesuchids is greater in the Karoo than anywhere else in Gondwana, and only in the Karoo are rhinesuchids known to have survived the end-Permian extinction event. These facts are significant because the Rhinesuchidae are widely regarded as the most basal stereospondyls (Milner 1990; Schoch & Milner 2000; Yates & Warren 2000). The Mesozoic sequence plays host to all 10 higher-level temnospondyl taxa known from the Karoo, distributed as follows. The Lystrosaurus Assemblage Zone contains seven higher-level taxa, the Amphibamidae, Tupilako- sauridae, Rhinesuchidae, Lydekkerinidae, Rhytido- steidae, Mastodonsauridae and Trematosauridae. The overlying Cynognathus Assemblage Zone hosts the last two of these taxa in addition to the Brachyopidae, Laidleriidae, and, possibly, the Rhytidosteidae. The now greater diversity of higher-level temnospondyl taxa in the Lystrosaurus biozone is noteworthy because of the previously held notion that the Cynognathus biozone contained a greater diversity of temnospondyls. This shift in diversity has occurred only recently, however, with the resurgence in studies of Beaufort Group temnospondyls. The remaining higher-level taxon, the Chigutisauridae, is at present known only from the Late Triassic–Early Jurassic Elliot Formation. Finally, it is particularly noteworthy that the Lystrosaurus Assemblage Zone plays host to a relict member of the dvinosaurian radiation (Thabanchuia), the last surviving member of the Dissorophoidea (Micropholis), and the only Triassic rhinesuchid (Broomistega). These facts, combined with the high diversity of rhinesuchids in the Permian of the Karoo, lend strong support to the hypothesis that the initial radiation of the Stereospondyli occurred in a ‘safe haven’ that was located in Gondwana (Yates & Warren 2000; Warren et al. 2000). The evidence for a Gondwanan origin and radiation also suggests that the exact position of the ‘safe haven’ may have been southern Africa. How- ever, it remains unclear whether the great stereospondyl ISSN 0078-8554 Palaeont. afr. (December 2003) 39: 21–36 33 radiation began in the Late Permian (Yates & Warren 2000) or in the earliest Triassic (Milner 1990). We are grateful to Mrs Elizabeth Latimer-Schaafsma (BPI Palaeontology) for generously sharing information on rhinesuchids, Dr Alain Renaut (BPI Palaeontol- ogy) for locality information, and Dr Andrew Milner (Birkbeck College, London) for information on Micropholis stowii and ‘Petrophryne major’. 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